AMPHIBIA: ANURA: LEPTODACTYLIDAE

Catalogue of American Amphibians and Reptiles.

Heyer, W.R. and M.M. Heyer. 2001. Leptodactylus lithonaetes.

Leptodactylus lithonaetes Heyer

Leytodachlus lithonaeres Heyer 1995:708. Type locality, "SW sector Cerro Yapacana, 600 m, 3'57'N, 67'00'W, Venezuela; Amazonas." Holotype, American Museum of Natural His- tory (AMNH) 100656, collected by Charles W. Myers and John Daly on 18-19 February 1978 (examined by WRH).

hptodach l~ is 1irhornere.s: La Marca 1997: 117. Lapsus. f.ep1odachlus lithonasres: Renjifo 199754. Lapsus.

CONTENT. This species is monotypic.

DEFINITION. Adult h p ~ o d a c h l u s lirhoncretes are of mod- erate size. the head length and head width are about equal. and the hind limbs are of moderate length (Table; for size defini- tions, see Heyer and Thompson 2000). Male vocal sacs are laterally expanded and darkly pigmented. Adult males have one black thumb spine on each hand and a pair of chest spines. Males have a patch of black tubercles on the chin and anterior portion of the throat. and lager males also have a band of black tubercles across the chest. Most individuals either lack dorso- lateral folds or exhibit one short pair of ridges or elongate warts in the shoi~lder region. Some individuals have series of warts or ridges in the dorsolateral fold field that extend the entire length of the dorsum from behind the eye or extend only to the sacrum.

The toe tips are bulbous and the toes are free of webbing and lateral fringes. The upper shank and outer tarsus have few to many black andlor white tubercles and may or may not have a shagreen (sharkskin-like surface). The sole of the foot is smooth in most specimens (86%), some (12%) have only a few black and/or white tubercles, and few individuals (2%) have a weakly developed shagreen. The upper lip pattern demonstrates a con- tinuum of uniform (13%). alternating light and dark vertical bars (32%), alternating light and dark oblique bars (29%), and an irregularly defined light area in the loreal region (26%). Most specimens have a dorsal pattern consisting of a series of 3 4 (rarely 2) pairs of spots ranging from small to large in size and from discrete to patterns of fusion with other spots both across and lengthwise along the dorsum; 10% of individuals have a uniform dorsum. The species lacks light middorsal stripes. The belly pattern ranges from uniformly gray-brown to boldly mottled with light spotslflecks on a darker ground. Belly pat- terns of iuveniles and adults do not differ markedly. The voste- nor thigh surface patterns demonstrate a continuum ranging from indistinctly mottled through distinctly mottled with small or large light irregular spots or light vertical bars on the upper thigh sur- face to a large light area on the upper thigh surface.

Larvae are elongate and depressed with low tail fins, and are members of the semiterrestrial guild (Altig and Johnson 1989). The larvae have a series of glandular ridges on the body above the abdominal cavity. The oral disk is ventrally positioned, emarginate, and with an anterior gap. The tooth row formulae are 2(2)/3 or 2(2)/3(1). The spiracle is sinistral and the anal tube is median. Total length ranges from28-36 nlm (for Gosner stage 3 5 4 0 larvae). For the same larvae, the head-body length ranges frotn 9-10 mm, the eye diameter is 14-16% of the head- body length, and the oral disk width is 22-26% of the head- body length. The dorsum of the head-body is brown, the under- side of the head-body is with or without white flecks whereas the region anterior to the guts has melanophores visible under the skin. The dorsal tail musculature and dorsal fin are mottled with or without darker flecks and/or light ocelli; the ventral tail musculature and ventral fin either lack melanophores or have melanophores on the distal half of the tail.

The advertisement call has not been recorded, but see De- scriptions.

DIAGNOSIS. Adult LRptodac~lus lithor~aetes have toes free of webbing and lateral fringes and indistinct dorsolateral folds. These features are shared with at least some individuals of L. bufonius, L. labicrlis, L. labyrinthicus, L. lnticeps, L. Iatirzosus,

FIGURE 1. Adult male Leptodact?.lus lirhonaetes (AMNH 100668) from the southwestern sector of Cerro Yapacana, Amazonas, Venezuela (photograph courtesy of Charles W. Myers, Americ'an Museum of Natural History).

FIGURE 2. Tadpole of Lrp~odrrcrylus lirhonaetes, UTA-A 23509, stage 38. TL 34.3 mm. Scale bar for larva 5 mm, for mouthparts I mm.

L. myersi, L. rugosus, L. syphax, and L. troglodytes. The upper shank and posterior tarsus of L. bufonius, L. labialis, L. larinasus, and L. troglodytes are covered with large prominent white tu- bercles and males lack thumb spines. In contrast, the upper shank and posterior tarsus of L. lithonaetes have black-tipped tubercles (in some preserved specimens the black tips may be lost, leaving white tubercles which are noticeably smaller than those of L. bufonius, etc.) and the males have a spine on each thumb. Leptodacfylus laticeps has a tile-like dorsal pattern and is larger (minimum adult SVL91 mm) than L. lithonaetes (maxi- mum adult SVL 78 m), which lacks a distinct tile-like pattern. Leptodac~ylus labyrinthicus is larger (minimum adult SVL 117) than L. lithonaetes, and no L. labyrinthicus have light loreal blotches, whereas several L. lithonaetes do. Lepfodnctylus myersi is larger (females 104-1 13 mm SVL, males 74-1 18 mm SVL) than L. lithonaetes, and L. myer.si males lack the chest spines found on L. lirhonaetes. Leptodactyhis lithonaeres is most likely to be confused with L. rugosus and L. syphax, but all demonstrate distinctive male secondary sexual characteristics: L. lithonaetes has a single black spine on each thumb and a patch of brownlblack tubercles on the chintthroat; all L. syphax and some L. rugosus have two spines on each thumb, and no L. syphax or L. rugosus males have a patch of chin tubercles.

Larvae lack dorsal fins on the anterior half of the tail, a con- dition shared only with L. rugosrrs larvae among congeners for

TABLE. Summary measurement data for Leprodacrylus lithonaetes. Means are given in parentheses.

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Measurement Males Females

SVL (mm) 45-71 (56.3) 55-78 (63.6) Head IengtWSVL (%) 37-44 (40) 3 M 3 (39) Head width/SVL (%) 3 8 4 2 (40) 3-1 (39) Thigh IengtWSVL (%) 3 7 4 8 (42) 3 9 4 5 (43) Shank IengtWSVL (%) 40-48 (43) 3 8 4 7 (44) Foot IengtWSVL (%) 42-53 (47) 40-51 (46)

which larvae have been described. The larvae of these two spe- cies are quite similar to each other and may be indistinguish- able. For the limited samples available, the most diagnostic feature is the number of labial teeth on one side of the split tooth row anterior to the beak (row A-2) for Gosner stage 3 3 4 2 larvae: 37-5 1 for L. lithoncretes and 50-56 for L. rugosus.

DESCRIPTIONS. Heyer (1995) listed morphological char- acteristics of the holotype and provided detailed character de- scriptions of adults, juveniles, and tadpoles, including color patterns. Rivero's (1961) data for Leptodac~ylus rugosus, in- cluding that pertaining to adult morphology, apparently all per- tain to L. lithonaetes (all the specimens he cited at the top of the species account are L. lithonaefes). He described the advertise-

MAP. Distribution of Leprodochlus lilhontietes; the circle marks the type locality, dots indicate other published records, and each x indicates an unpublished record. A symbol may cover more than a single locality. The question mark represents a locality (Cerro Guanay, Bolivar. Venezuela) for which species identification is in doubt (see Distribution). Because the species occurs in rocky outcrops that are patchily dislributed throughout its distributional range, a predictive range outline would be misleading.

ment call as "very similar to the noise produced when a stone is thrown into a quiet pool of water." We know of no recordings of the call of L. lithonnetes. Rivero's description is very differ- ent from the call described by Lynch and Vargas (2000) as "una silbato mecinico, colno un juguete de cuerda" (a mechanical whistle, like a string toy). Rivero's description probably ap- plies to L. sabanensis or one of its close relatives.

LLLUSTRATIONS. Color photographs of adult males are in Heyer ( 1995) and Renjifo (1997). Pictures ofjuveniles and draw- ings of tadpoles are in Heyer ( 1995). Rada de Martinez (1990) illustrated tadpoles identified as L. ritgosus, but we were unable to find the locality (Pozo Pio, T. F. Amazonas, Venezuela) to determine whether the larvae are L. lithonnetes or L. rugosus.

DISTRLBUTION. Leptodac~ylus 1ithotrnete.s is known from a few localities near the border between Colombia and Venezu- ela and extending eastward into Colombia. The species appar- ently is restricted to granitic and sandstone habitats associated with flowing water. Duellman (1999) tabled its distribution as Amazon Basin-Guiana lowlands. A range map was provided by Heyer (1995).

The Venezuelan locality of Cerro Guanay, Bolivar (Gorzula and Seiiaris 1999) lies between the distributions of L. lithonnetes and L. rugosus as mapped by Heyer (1995). These specimens are juveniles; adult males need to be examined from that local- ity to determine whether the specimens are L. 1ithotraete.s or L. rugosus. The specimens of L. rugosus reported by Parker (1936) and collected by the Marquis de Wavrin from the upper Orinoco in the state of Amazonas, Venezuela are probably L. lithonrtetes.

FOSSrL RECORD. None.

PERTINENT LITERATURE. Relatively little literature concerns Leptod(~c~yhts 1ithotrcrete.s. Rivero ( I96 1 ) reported on the natural history of L. lithotrctetes (as L. rugosus). Skin com- pounds of L. 1ithonerete.s specimens were tabled as L. rrcgosus in Flier et al. (1980; see Heyer and Thompson 2000 for docutnen- tation of identification). C.W. Myers (in Heyer 1995) described habitat and behavior. Relationships were analyzed in Heyer (1995) and Eterovick and Sazima (2000). Barrio (1998) pro- vided a brief species account. Barrio (1998) and La Marca (1997) included L. Iithonaetes in faunal lists for Venezuela, and Glaw et al. (1998) cited it in a list of new species. Many literature references to L. rugosus prior to 1995 and some citations after that date are unclear as to whether they refer to L. lithonoetes, L. nrpersi, or L. rugosus (see Heyer and Thompson 2000).

REMARKS. Barrio (1998) coined a rather unwieldy com- mon name for Leptode~crylus 1ithonarte.s. sapo-rana rugoso occidental. Because we believe that colnlnon names should be designated by the people living in the region where the species occurs, and we are unaware of any such name for L. lithonnetes. we propose that the species should be referred to by its scien- tific name for all purposes.

ETYMOLOGY. Heyer (1995) derived the name lirhortneres from the Greek lithos, stone or rock, and naere.7, inhabitant, in reference to the habitat of this species.

*ACKNOWLEDGMENTS. John D. Lynch reviewed the pa- per and provided assistance with localities. Cesar Luis Barrio

Amarbs clarified certain localities and provided new, unpub- lished records for Venezuela. P. E. Vanzolini also provided help with problematic localities. Preparation of this account was supported in part by the Neotropical Lowlands Research Pro- gram, Smithsonian Institution (Richard P. Vari, Principal Inves- tigator) and the National Science Foundation (award 9815787 to Rafael de SA and WRH).

LITERATURE CITED

Altig, R. and G.F. Johnston. 1989. Guilds of anuran larvae: relation- ships among developmental modes, morphologies. and habitats. Her- petological Monographs 3:8 1-1 09.

Barrio, C. 1998. Sistemstica y biogeograria de 10s anfibios (Amphibia) de Venezuela. Systematics and biogeography of the amphibians (Am- phibia) of Venezuela. Acta Biologica Venezuelica 18: 1-93.

Duellman, W.E. 1999. Distribution patterns of amphibians in South America, p. 255-328. In W.E. Duellman (ed.), Patterns of Distribu- tion of Amphibians. A Global Perspective. The Johns Hopkins Uni- versity Press. Baltimore and London.

Eterovick. P.C. and I. Sazima. 2000. Description of the tadpole of Leprodacn.lus sjpliax, with a comparison of morphological and eco- logical characters of tadpoles and adults of the species in the L. pentodac~lus group (Leptodactylidae. Anura). Aniphibia-Reptilia 2 1 :34 1-350.

Flier, J., M.E. Edwards, J.W. Daly, and C.W. Myers. 1980. Widespread occurrence in frogs and toads of skin compounds interacting with the ouabian site of Na', K'-ATPase. Science 208503-505.

Gorzula, S. and J.C. Sefiaris. 1999. Contribution to the herpetofauna of the Venezuelan Guayana I. A data base. Scientia Guaianae (8):xviii + 269 p. + 32 color plates.

Heyer. W.R. 1995. South American rocky habitat Leprodarhlus (Am- phibia: Anura: Leptodactylidae) with description of two new spe- cies. Proceedings of the Biological Society of Washington 108:695- 716.

- and A.S. Thon~pson. 2000. Lepro(locn1trs rirgosrrs. Catalogue of American Amphibians and Reptiles (708): 1-5.

La Marca. E. 1997. Listn actualizada de 10s anfibios de Venezuela, p. 103-120. In E. La Merca (ed.). Vertebrados Actuales y F6siles de Venezuela. Serie CatAogo ZoolBgico de Venezuela. Vol. I . Museo de Cienci;~ y Tecnologh de Merida. Merida. Venezuela.

Lynch. J. D. and M. A. Vargas. 2000. Lista preliminar de especies de anuros del Departamento del Guainia, Colombia. Revista de la Aca- demia Colombiana de Ciencias Exactas. Fisicas y Naturales 24579- 589.

Parker, H.W. 1936. A collection of reptiles and amphibians from the Upper Orinoco. Bulletin du Musee royal d'Histoire naturelle de Belgique 12: 1 4 .

Rada de Maninez. D. 1990. Contribution al conocimiento de las larvas de anfibios de Venezuela. Memoria, Sociedad de Ciencias Naturales La Salle 49-50:391403.

Renjifo, J.M. 1997. Ranas y Sapos de Colombia. Instituto Humboldt. Colombia.

Rivero. J.A. 1961. Salientin of Venezuela. Bulletin of the Museum of Comparative Zoology 126: 1-207.

W. RONALD HEYER and MIRIAM MUEDEKING HEYER, Amphibians & Reptiles, MRC 162, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0 162, USA.

Primary editor for this account, Hinrich Kaiser.

Published 30 June 2001 and Copyright O 2001 by the Society for the Study of Amphibians and Reptiles.